Effects of clipping and soil compaction on growth,morphology and mycorrhizal colonization of Schizachyrium scoparium,a C4 bunchgrass

Summary A factorial design of clipping and compaction was used to study the responses of Schizachyrium scoparium and its mycorrhizal symbionts to these stresses. All treatment combinations significantly reduced the growth and biomass of plants relative to controls. Compaction significantly reduced tillering and crown expansion while clipping increased tillering early in the growing season and reduced it later. Mycorrhizal colonization of roots was highest in the clipped plots and lowest in compacted plots. Spore number was highest in compacted plots and lowest in clipped plots. It appears that spore number may be negatively correlated with root growth since any treatment that reduced plant growth yielded higher spore numbers. The combination of clipping and compaction reduced plant growth the most, but had intermediate effects on mycorrhizal colonization and spore number.     

Effects of endomycorrhizal infection,artificial herbivory,and parental cross on growth of Lotus corniculatus L.

Summary We examined how combinations of parentage, fungicide application, and artificial herbivory influence growth and shoot phosphorus content in pre-reproductive Lotus corniculatus, using young offspring arising from three parental crosses, two of which had one parent in common. Soil with vesicular-arbuscular mycorrhizal (VAM) fungi was treated with either water or benomyl, an anti-VAM fungicide, and added to trays containing groups of four full siblings. There were two experiments; in the first no plants were clipped while in the second two of the four plants were clipped to simulate herbivory. In both experiments plants of the two related crosses accumulated more biomass and total shoot P than did plants of the third cross. Plants inoculated with watertreated soil had greater shoot mass and P concentration than did fungicide-treated replicates but the extent of increase in P concentration varied among crosses. In Experiment 2, clipping reduced root mass and resulted in higher shoot P concentration. In this experiment there was a significant interaction of fungicide application and clipping: both unclipped and clipped plants grew better in soil not treated with fungicide, but the increase in shoot mass, total mass, and total P was greater in unclipped plants. Significant interaction of fungicide treatment and clipping is most likely due to reduced availability of carbon to the roots of clipped plants, resulting in poorer symbiotic functioning.     

GROWTH OF LITTLE BLUESTEM (SCHIZACHYRIUM SCOPARIUM) (POACEAE) IN FUMIGATED AND NONFUMIGATED SOILS UNDER VARIOUS INORGANIC NUTRIENT CONDITIONS

Little bluestem plants (Schizachyrium scoparium (Michx.) Nash) were grown in fumigated and nonfumigated soil under manipulated levels of three inorganic nutrients: nitrogen, phosphorus, or bases (Ca + Mg). Plants grown in nonfumigated soil had significantly (P < 0.05) higher tissue levels of inorganic nutrients (Cu, Zn, Al, S, Mg, Mn, Ca, and P), smaller shoots, less total biomass, fewer flowering plants but more VAM fungal colonization than plants grown in fumigated soil that were essentially nonmycorrhizal (colonization vs. 1.2 ± 4.9%, for plants grown in nonfumigated and fumigated soil, respectively). Levels of phosphorus (14–33 μg/g) available (Bray No. 1) in the soil prior to manipulation, which are adequate for little bluestem, likely resulted in the development of an ineffectual mycorrhizal association, which in turn, caused the depressed growth of plants in nonfumigated soil. Among plants grown in nonfumigated soil, there was significant variation in VAM fungal colonization and sporulation owing to nutrient treatment. Nitrogen treatment and deionized water control had significantly lower levels of colonization than phosphorus and base treatments. However, plants in the nitrogen and base treatments had significantly more spores/100 cc of rhizosphere soil than plants grown in the deionized water control.     

A test of compensatory photosynthesis in the field: Implications for herbivory tolerance

Summary The occurrence of compensatory photosynthesis was examined in the field for all foliage elements on two Agropyron bunchgrass species that differ in their evolutionary history of grazing pressure. This is the first reported field study of compensatory photosynthesis in individual foliage elements of graminoids. Compensatory photosynthesis was defined as an increase in the photosynthetic rates of foliage on partially defoliated plants relative to foliage of the same age on undefoliated plants. Compensatory photosynthesis did occur in many individual foliage elements during at least part of their ontogeny. For both species, compensatory photosynthesis was related primarily to delayed leaf senescence and increased soluble protein concentrations, but not to an improvement in the water status of clipped plants. Soluble protein concentration increased in all foliage elements. A delay in senescence on clipped plants was documented for the two oldest, fully-expanded leaves that were present when the plants were initially clipped, but the initiation and senescence of all other foliage elements were not affected by the clipping treatments. Photosynthetic water use efficiency and photosynthetic rates per unit soluble protein of foliage on partially defoliated plants were not increased following the clipping treatments. Although A. desertorum and A. spicatum were exposed to different levels of grazing pressure during their evolutionary history, the phenology, water status, and gas exchange rates of foliage were very similar both for undefoliated as well as partially defoliated plants. Thus, we conclude that compensatory photosynthesis does not appear to be an important ecological component of herbivory tolerance for these species.     

Compensatory growth and photosynthetic responses of <Emphasis Type="Italic">Pharbitis purpurea</Emphasis> seedlings to clipped cotyledon and second leaf

Leaf tissue damaging to seedlings can limit their subsequent growth, and the effects may be more extensive. Compensatory photosynthesis responses of the remnant cotyledon and primary leaf of Pharbitis purpurea to clipping and the effect of clipping on seedling growth were evaluated in a pot-cultivated experiment. Three treatments were conducted in the experiment, which were clipped cotyledon (CC), clipped second leaf (CL), and control group (CG). The area, thickness, mass, and longevity of the remaining cotyledon of CC exhibited over-compensatory growth. In contrast, seedlings of CC had under-compensatory growth in seedling height, root length, seedling mass, and root to shoot ratio. However, the traits of remnant cotyledon and seedling in CL treatment exhibited equal-compensatory growth. Net photosynthetic rate of the cotyledon of CC was significantly higher than those of CL and CG treatments, and the diurnal changes in photosynthetic rates showed significantly different patterns which were unimodal curve (CC) and bimodal curve (CL and CG), respectively. There was no significant difference between CL and CG treatment. Net photosynthetic rate of the primary leaf of CL was significantly higher than that of CG treatment. However, the photosynthetic rates of primary leaves of CL and CG treatments showed similar photosynthetic patterns characterized by a bimodal curve. P. purpurea seedlings used a compensatory growth strategy in the remaining cotyledon or the primary leaf to resist leaf loss and minimize any adverse effects.     

Compensatory growth responses to clipping defoliation in Leymus chinensis (Poaceae) under nutrient addition and water deficiency conditions

Compensatory growth responses of Leymus chinensis, a dominant species in Inner Mongolia steppe, to clipping defoliation were evaluated in a pot-cultivated experiment under different nutrient (N and P) and water availability conditions. Leymus chinensis exhibited over-compensatory growth at the light and moderate clipping intensities (20% and 40% aerial mass removed) with a greater accumulated aboveground biomass, higher relative growth rate (RGR), more rhizomatic tillers and a stimulation of compensatory photosynthesis to the remnant leaves as compared with those of the unclipped plants. Intense clipping (80% aerial mass removed), which removed most of the aboveground tissues, greatly reduced the growth of aboveground biomass in comparison with that of the unclipped plants. Nitrogen addition only slightly improved the biomass production and RGR in light and moderately clipped plants, and it did not allow plants in the intense clipping condition to over-compensate. Phosphorus addition had no obvious influences on the growth and physiological responses to clipping defoliation. These results indicated that nutrient addition could not compensate for the negative effects of severe clipping on the defoliated grass. On the other hand, there were no distinct positive responses under water deficiency condition for L. chinensis at all clipping intensities with a significant reduction of aboveground and belowground biomass, lower RGR, fewer rhizomatic tillers, and a lower net photosynthetic rate than other wet treatments. Additionally, the chlorophyll contents of remnant leaves gradually increased with the increase of clipping intensities in each treatment. In conclusion, although L. chinensis could compensate for tissues removal by some morphological and physiological responses, intense clipping and drought can result in a significant decrease of biomass and growth rate, even under enriched nutrition conditions.     

Biomass and mineral element responses of a Serengeti short-grass species to nitrogen supply and defoliation: compensation requires a critical [N]

Large mammalian herbivores in grassland ecosystems influence plant growth dynamics in many ways, including the removal of plant biomass and the return of nutrients to the soil. A 10-week growth chamber experiment examined the responses of Sporobolus kentrophyllus from the heavily grazed short-grass plains of Serengeti National Park, Tanzania, to simulated grazing and varying nitrogen nutrition. Plants were subjected to two clipping treatments (clipped and unclipped) and five nitrogen levels (weekly applications at levels equivalent to 0, 1, 5, 10, and 40 g N m⁻²), the highest being equivalent to a urine hit. Tiller and stolon production were measured weekly. Total biomass at harvest was partitioned by plant organ and analyzed for nitrogen and mineral element composition. Tiller and stolon production reached a peak at 3–5 weeks in unclipped plants, then declined drastically, but tiller number increased continually in clipped plants; this differential effect was enhanced at higher N levels. Total plant production increased substantially with N supply, was dominated by aboveground production, and was similar in clipped and unclipped plants, except at high nitrogen levels where clipped plants produced more. Much of the standing biomass of unclipped plants was standing dead and stem; most of the standing biomass of clipped plants was live leaf with clipped plants having significantly more leaf than unclipped plants. However, leaf nitrogen was stimulated by clipping only in plants receiving levels of N application above 1 g N m⁻² which corresponded to a tissue concentration of 2.5% N. Leaf N concentration was lower in unclipped plants and increased with level of N. Aboveground N and mineral concentrations were consistently greater than belowground levels and while clipping commonly promoted aboveground concentrations, it generally diminished those belowground. In general, clipped plants exhibited increased leaf elemental concentrations of K, P, and Mg. Concentrations of B, Ca, K, Mg, and Zn increased with the level of N. No evidence was found that the much greater growth associated with higher N levels diminished the concentration of any other nutrient and that clipping coupled with N fertilization increased the total mineral content available in leaf tissue. The results suggest that plants can (1) compensate for leaf removal, but only when N is above a critical point (tissue [N] 2.8%) and (2) grazing coupled with N fertilization can increase the quality and quantity of tissue available for herbivore removal. Received: 25 August 1997 / Accepted: 14 April 1998     

Response of two prairie forbs to repeated vole herbivory

Vertebrate herbivores as diverse as ungulates, geese, and rabbits preferentially feed on plants that have previously experienced herbivory. Here, we ask whether smaller grassland “cryptic consumers” such as voles (Microtus ochrogaster and M. pennsylvanicus) preferentially clip (cut stems for access to leaves or seeds) or avoid previously clipped individuals of two tallgrass prairie species (Desmanthus illinoensis and Echinacea purpurea) within a growing season. Further, we ask how these plants respond to repeated clipping within a growing season, and whether the effects of this herbivory last into the subsequent growing season. Voles preferentially clipped stems of D. illinoensis and E. purpurea plants that had been previously clipped. The exception was indiscriminant clipping of stems of E. purpurea late in the growing season when its achenes, a favorite vole food, ripened. For D. illinoensis, repeated clipping resulted in a 59% reduction in biomass, 42% lower ratio of reproductive to vegetative biomass, and 57% fewer seeds produced per plant compared with unclipped plants. These effects lasted into the following growing season in which plants were protected from voles. In contrast, the only effect of repeated clipping for E. purpurea was that the number of achenes per plant was substantially reduced by three episodes of clipping. This effect did not carry over to the next growing season. Such differences in D. illinoensis and E. purpurea response to repeated stem clipping by voles offer insights into how these small rodents can effect major changes in composition and dominance in grassland communities.     

Root growth response to defoliation in two Agropyron bunchgrasses: field observations with an improved root periscope

Summary Root growth responses to defoliation were observed in the field with an improved root periscope technique, which is described. The grazing tolerant, Eurasian bunchgrass, Agropyron desertorum, was compared with the very similar but grazing sensitive, North American bunchgrass, A. spicatum. Root length growth of clipped A. desertorum was about 50% of that of intact plants, while root elongation of clipped A. spicatum continued relatively unabated during ninety days of regrowth following severe defoliation. The reduced root growth in A. desertorum was correlated with the allocation of relatively more resources to aboveground regrowth, thus aiding reestablishment of the root: shoot balance. This balance was apparent in similar root mortality patterns of clipped and control A. desertorum plants in the season following defoliation. In clipped A. spicatum, however, root mortality increased in the winter following the season in which the clipping was done and continued into the subsequent growing season. Reduction of root growth following defoliation appears to be an effective mechanism to aid reestablishment of the photosynthetic canopy and the root: shoot balance. As such it contributes to both herbivory tolerance and maintenance of competitive ability.     

Leaf compensatory growth as a tolerance strategy to resist herbivory in Pancratium sickenbergeri

We examined the effects of leaf herbivory by the dorcas gazelle, Gazella dorcas, on the compensatory growth of the geophyte Pancratium sickenbergeri (Amaryllidaceae) in the Negev desert, Israel. In three populations exposed to different levels of herbivory, we removed different amounts of photosynthetic leaf area from plants in five clipping treatments: 0, 25, 50%-dispersed over all leaves, 50%-entire area of half the leaves, 100%. The population with the lowest level of herbivory showed the lowest relative regrowth rate after clipping. In the population with a constantly high level of herbivory, plants in intermediate-clipping treatments overcompensated in leaf area after clipping. For all the populations, clipped plants produce more new leaves than unclipped plants. In the population with the highest level of herbivory, clipping treatments did not have a significant effect on the number of fruits per plant. In addition, we did not find a trade-off between investments in growth and reproduction in this population. Our results indicated that, in the desert lily, herbivores may select for plant mechanisms that compensate after damage as a tolerant strategy to maintain fitness.     

The Effect of Clipping on Net Photosynthesis and Dark Respiration Rates of Plants from an Upland Grassland, with Reference to Carbon Partitioning in Festuca ovina

Four co-existing species (Deschampsia flexuosa, Festuca ovina,Juncus squarrosus and Nardus stricta) were subjected to clippingand the net photosynthetic and dark respiration rates were followedafter this treatment for 50 d. Concurrently carbon partitioningin F. ovina plants clipped initially and again at 50 and 100d was examined. An expansion of new leaf lamina was observed with F. ovina,which had a greater net photsynthetic rate per unit leaf areathan unclipped lamina. The remaining leaf lamina (stubble) afterclipping also showed net photosynthetic and dark respirationrates greater than unclipped lamina; these responses were uniqueto F. ovina plants. N. stricta was the only other species toattain a pre-clipping photosynthetic rate within 6 d. Clipped F. ovina plants showed a change in carbon allocationpattern, with a reduction in carbon allocated to roots. ¹⁴Caccumulated in roots and stubble was shown to have a role inregrowth, as was current assimilate via the production of newleaf lamina. Plants initially clipped before exposure to ¹⁴C,redistributed less ¹⁴C to new shoot growth and, therefore, lostless when subsequently clipped. Further redistribution of ¹⁴Ccame from leaf stubble tissue and not at the expense of roots.The variation between species in clipping response are discussedin terms of the implications for coexistence. Carbon partitioning, clipping, gas exchange, grasses     

Growth,morphology and gas exchange of mycorrhizal and nonmycorrhizal Panicum coloratum L., a C4 grass species,under different clipping and fertilization regimes

Summary Root samples collected in grasslands of the Serengeti ecosystem, Tanzania, were found to be mycorrhizal and infection frequency was positively correlated with grazing intensity across sites. To examine the role of mycorrhizae in a grazing ecosystem, I analyzed the growth, morphology and gas exchange of mycorrhizal and nomycorrhizal plants of Panicum coloratum L. under different fertilization and clipping regimes. Both severe clipping and high nitrogen promoted more prostrate shoot growth but inhibited root growth. However, mycorrhizal infection promoted a prostrate shoot morphology and enhanced root growth. Photosynthesis was inhibited by clipping, however; at the most severe clipping and nitrogen regime, photosynthesis of the mycorrhizal plants was not affected whereas the largest inhibition of photosynthesis occurred in similarly treated nonmycorrhizal plants. Discussion of the putative roles of mycorrhizae in intensely grazed ecosystems is presented.     

Responses of an African graminoid (Themeda triandra Forsk.) to frequent defoliation,nitrogen, and water: a limit of adaptation to herbivory

Summary Themeda triandra Forsk., a medium height perennial grass common in semi-arid Africa, was collected from Serengeti National Park in Tanzania, propagated vegetatively and grown in controlled environments simulating native conditions. The experiment demonstrated mechanisms that result in inability to withstand frequent defoliation, and thus how plant-herbivore interactions are affected by plant morpho-physiology. Clipping reduced most plant yield components. After two months leaf elongation rate was greater in clipped plants, but over the whole experiment aboveground yield was unaffected by clipping. Since clipping did not stimulate growth, residual leaf area was less in clipped plants. Tillering was not stimulated by clipping so there was no mechanism to increase leaf area below the clipping height. The resulting reduced leaf area accounted for reduced root and crown production by clipped plants.     

Seasonality of herbivory and communication between individuals of sagebrush

Seasonal changes in herbivore numbers and in plant defenses are well known to influence plant–herbivore interactions. Some plant defenses are induced in response to herbivore attack or cues correlated with risk of attack although seasonal variation in these defenses is relatively poorly known. We previously reported that sagebrush becomes more resistant to its herbivores when neighboring plants have been experimentally clipped with scissors. In this study we asked whether herbivory to leaves of sagebrush varied seasonally and whether there was seasonal variation in natural levels of damage when neighbors were clipped. We found that sagebrush accumulated most chewing damage early in the season, soon after the spring flush of new leaves. This damage was caused by generalist grasshoppers, deer, specialist caterpillars, beetles, gall makers, and other less common herbivores. Sagebrush showed no evidence of preferentially abscising leaves that had been experimentally clipped. Experimental clipping by Trirhabda pilosa beetle larvae caused neighbors to accumulate less herbivore damage later that season, similar to results in which clipping was done with scissors. Induced resistance caused by experimentally clipping a neighbor was affected by season; plants with neighbors clipped in May accumulated less damage throughout the season relative to plants with unclipped neighbors or neighbors clipped later in the summer. We found a correlation between seasonal herbivore pressure, damage accumulated by plants, and induced responses to experimentally clipping neighbors. The causal mechanisms responsible for this correlation are unknown although a strong seasonal effect was clear.     

Does pollination limit tolerance to browsing in <Emphasis Type="Italic">Ipomopsis aggregata</Emphasis>?

Ungulate browsing of flowering stalks of the semelparous herb Ipomopsis aggregata leads to regrowth of lateral inflorescences, a response that has been reported to yield overcompensation in some cases (browsed plants with higher reproductive success than unbrowsed), but undercompensation in others. Little is known about the mechanisms that cause such variable tolerance to herbivory. We explored one possible mechanism—variation in effects of browsing on pollination—by clipping I. aggregata inflorescences to mimic browsing, observing subsequent visits by pollinators and nectar-robbers, and adding pollen by hand to flowers of some clipped and unclipped plants. Clipping reduced floral display size and increased inflorescence branching, but neither hummingbirds, the primary pollinators, nor nectar-robbing bumblebees showed any preference for unclipped versus clipped plants. Clipping delayed flowering; this shift in phenology caused clipped plants to miss the peak of hummingbird activity and to have lower per-flower visitation rates than unclipped controls in one year, but to have greater overlap with birds and higher visitation rates in the subsequent year. In three sites and 2 years, clipped plants exposed to natural pollination suffered extreme undercompensation, producing on average only 16% as many seeds as unclipped controls. This was not directly attributable to clipping effects on pollination, however, because clipped plants were unable to increase fecundity when provided with supplemental pollen by hand. Taken altogether, our results suggest that compensation was constrained less by indirect effects of browsing on pollination than by its direct impacts on resource availability and hence on the ability of plants to regrow lost inflorescence tissue and to fill seeds. Exploring the physiological and developmental processes involved in regrowth of inflorescences and provisioning of seeds is a promising future direction for research designed to understand variation in browsing tolerance.     

The interaction of defoliation and nutrient uptake in Sporobolus kentrophyllus,a short-grass species from the serengeti plains

Summary Sporobolus kentrophyllus, a grazing-tolerant C₄ grass from the southeastern Serengeti Plains, was grown in solution culture to examine the effects of clipping on the uptake, preference and subsequent transport of varying nitrogen forms. Clipping reduced offtake mass, crown mass ane root mass, resulting in a 58% decline in plant mass. Proportional biomass allocation to roots decreased with clipping, while tillering rates increased. Clipping also increased the nitrogen concentrations of all tissues, and plant nitrogen uptake (nitrogen accumulated throughout the experiment per gram root). The ¹⁵N concentrations (% atom excess) of all tissues were higher in clipped compared with unclipped plants, and the average ¹⁵N uptake rate of clipped plants was twice that of unclipped plants. The relative ¹⁵N allocation to aboveground mass, a measure of canopy sink strength, was higher in clipped plants. Plants fed ¹⁵N-ammonium or ¹⁵N-nitrate during the ¹⁵N pulse experiment had greater ¹⁵N tissue concentrations compared with urea-fed plants, and ¹⁵N uptake rates were higher in ammonium-fed and nitrate-fed plants, compared with urea-fed plants. The relative magnitudes of these differences were higher when plants were clipped. Clipped plants had higher uptake rates for potassium, phosphorus and sodium, while differences between clipping treatments for calcium, iron, and magnesium were indistinguishable. Rapid uptake rates for species on the southeastern Serengeti plains, particularly during grazing periods, have important implications for nutrient cycling in this system.     

Effects of simulated winter browsing on mountain birch foliar chemistry and on the performance of insect herbivores

Winter browsing by mammalian herbivores is known to induce a variety of morphological and physiological changes in plants. Browsing has been suggested to decrease the carbohydrate reserves in woody plants, which might lead to reduced tannin production in leaves during the following summer, and consequently, to increased herbivore damage on leaves. We conducted a clipping experiment with mature mountain birch trees and measured the effects of clipping on birch growth, leaf chemistry and toughness, as well as on the performance of insect herbivores. Leaves grew larger and heavier per unit area in the clipped ramets and had a higher content of proteins than leaves in the control trees. Clipping treatment did not affect the total content of sugars in the leaves (mg g^?1), suggesting that a moderate level of clipping did not significantly reduce the carbohydrate pools of fully‐grown mountain birch trees. Furthermore, the contents of proanthocyanidins (condensed tannins) and gallotannins were slightly higher in the leaves of clipped ramets, contrary to the hypothesis of reduced tannin production. The effects of clipping treatment on leaf and shoot growth and on foliar chemistry were mainly restricted to the clipped ramets, without spreading to untreated ramets within the same tree individual. The effects of clipping on leaf characters varied during the growing season; for instance, leaf toughness in clipped ramets was higher than toughness in control trees and ramets only when leaves were mature. Accordingly, clipping had inconsistent effects on insect herbivores feeding at different times of the growing season. The generally small impact of clipping on herbivore performance suggests that the low intensity of natural browsing at the study area, simulated by our clipping treatment, does not have strong consequences for the population dynamics of insect herbivores on mountain birch via enhanced population growth caused by browsing‐induced changes in food quality.     

Interactive regulation of grass yield and chemical properties by defoliation,a salivary chemical,and inorganic nutrition

Summary Clones of 2 C₄ grass species, Sprobolus ioclados and S. pyramidalis, were obtained from more and less heavily grazed grasslands, respectively, in Tanzania's Serengeti National Park. Plants were grown in a factorial experiment to determine the effects of severe defoliation, nutrient limitation, and a salivary chemical (thiamine) on plant growth, nitrogen content, and non-structural carbohydrate content. The experimental design included: (1) species; (2) clipping, with plants either unclipped or clipped weekly to a height of 5 cm; (3) 0.2 ml of distilled water of 0.2 ml of 10 ppb aqueous thiamine sprayed on plants from an atomizer after clipping and identical treatments at the same time to unclipped plants; (4) phosphorus (P) at 0.2 or 1 mM; (5) nitrogen (N) at 3 or 15 mM. Clipping was the major variable affecting plant growth. Total and litter yields were reduced to half and residual plant yield was reduced to 30% of the values for unclipped plants. Clipping interacted strongly with other variables since they commonly had minor effects on clipped plants and major effects on unclipped plants. Exceptions to this generalization were generally due to better performance by S. ioclados under clipping. Compared to lower treatment levels, higher treatment levels promoted total yield of unclipped plants by 52% for N, 43% for thiamine, and 33% for P. In general, thiamine had greater effects than P but lesser effects than N. Thiamine promoted yield and modified the chemical balance of plants by promoting carbohydrate (CHO) concentrations and reducing N concentrations. N and P deficiencies promoted CHO accumulation. Clipping promoted the N of leaves and crowns and reduced the N levels in roots. Leaf blade water and N contents were positively correlated with very little scatter. The slope of the line was different for S. ioclados and S. pyramidalis. Leaf blade water and CHO contents were negatively related but there was more scatter and the species could not be distinguished. The species from more heavily grazed grasslands was conspicuously more sensitive to thiamine application. The results indicated that leaf treatment with thiamine, the only likely source of which in natural grasslands is saliva deposited by feeding herbivores, can have major effects on plant yield and metabolic balances at very low application levels. But under defoliation levels as severe as those imposed in this experiment, which reduced above ground plant biomass to a fourth of the level produced by unclipped plants, growth was so strongly limited by defoliation that neither thiamine nor inorganic nutrients affected plant yield residual from clipping. Therefore, whether chemicals such as thiamine that may be introduced onto grass foliage by grazing ungulates and other herbivores will influence the growth of grazed plants will depend upon the grazing intensity associated with the transfer.     

Clipping at early florescence is more efficient for controlling the invasive plant <Emphasis Type="Italic">Spartina alterniflora</Emphasis>

As invasive plants in different life stages have different recovery potential against devegetation operation, how to optimize the proper timing (the time for single treatment or the starting time for repeated treatments) for eliminating them becomes an important topic in the management of invasive plants. The invasive plant Spartina alterniflora was selected in this study, and it was clipped as a devegetation example to examine how important treatment timing is for increasing control efficiency. The results illustrated that by either single or repeated clipping treatments within 1 year, clipping at florescence was more efficient for controlling S. alterniflora. On the other hand, increasing clipping frequency without proper timing might not enhance control efficiency, suggesting that treatments in proper timing might reduce the clipping frequency required for a specific control target. From a management point of view, each control method has its optimal treatment timing, thus, in order to improve the control efficiency, the control programs should be ideally designed in relation to the phenology of the target plant. Besides, repeated clipping showed significantly higher control efficiency in low tidal zone than in high tidal zone, implying that habitat dependence of invasive plant control efficiency is an important issue in the management of invasive plants.     

Responses of an African tall-grass (Hyparrhenia filipendula stapf.) to defoliation and limitations of water and nitrogen

Summary Hyparrhenia filipendula stapf., a tall (1–1.5 m) perennial grass common in dry-subhumid African savannas, was collected from Serengeti National Park in Tanzania, propagated vegetatively, and grown in controlled environments simulating conditions in nature. Plants were subjected to a factorial experiment with combinations of watering frequency, nitrogen supply, clipping height (10 and 15 cm) and clipping frequency (7 and 14 d). Biomass yield and allocation to various tissue types, morphometric traits, and growth processes were measured.Watering frequency affected leaf elongation rate while nitrogen affected tiller number. Clipped yield was strongly correlated with leaf elongation rate but not tiller number, therefore it was primarily controlled by the activities of intercalary rather than apical meristems. There was a negative exponential relationship between tillering and clipped yield per tiller. Plants that received both high nitrogen and high water closely followed a-3/2 power law in this tradeoff. The fraction of total net photosynthate allocated to roots was not significantly related to any environmental treatment. Root and crown growths were not affected by defoliation treatment; leaf blade and stem growths were inhibited; and sheaths were inhibited only under low water.Despite a tall stature, H. filipendula tolerated herbivory by increased photosynthetic rate (Wallace et al. 1984), through continued production of young tissues by intercalary meristems balanced against tiller number, and by a statistically constant proportional allocation to roots. Clipped yield increased only when both nitrogen and water were abundant, and then, proportional clipped yield did not surpass an upper asymptotic limit.